biology - pccua...respiration. • if o 2 is not available to the cell, fermentation, an anaerobic...

11

Chapter 8Cellular Respiration

Lecture Outline

BiologySylvia S. Mader

Michael Windelspecht

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Copyright © McGraw-Hill Education. Permission required for reproduction or display.

Outline• 8.1 Overview of Cellular Respiration• 8.2 Outside the Mitochondria:

Glycolysis• 8.3 Outside the Mitochondria:

Fermentation• 8.4 Inside the Mitochondria• 8.5 Metabolism

2

8.1 Cellular Respiration• Cellular respiration is a cellular process that breaks down

nutrient molecules produced by photosynthesizers with the concomitant production of ATP.

• It consumes oxygen and produces carbon dioxide (CO2). Cellular respiration is an aerobic process.

• It usually involves the complete breakdown of glucose to CO2and H2O. Energy is extracted from the glucose molecule:

• Released step-wise

• Allows ATP to be produced efficiently

Oxidation-reduction enzymes include NAD+ and FAD as coenzymes.

3

Cellular Respiration

4

Electrons are removed from substrates and received by oxygen, which combines with H+ to become water.

Glucose is oxidized and O2 is reduced.

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

energyglucose

Reduction

Oxidation

6H2O6CO26O2C6H12O6 + ++

Cellular Respiration

5


O2 and glucose enter cells,which release H2O and CO.

CO2

H2O

Mitochondria useenergy fromglucose to form ATPfrom ADP + P .

ATPADP + P

cristae

intermembranespace

© E. & P. Bauer/zefa/Corbis

6

Cellular Respiration• NAD+ (nicotinamide adenine dinucleotide) A coenzyme of oxidation-reduction, it is:

• Oxidized when it gives up electrons• Reduced when it accepts electrons

Each NAD+ molecule used over and over again

• FAD (flavin adenine dinucleotide) Also a coenzyme of oxidation-reduction Sometimes used instead of NAD+

Accepts two electrons and two hydrogen ions (H+) to become FADH2

Phases of Cellular Respiration

• Cellular respiration includes four phases: Glycolysis is the breakdown of glucose into

two molecules of pyruvate. • It occurs in the cytoplasm.• ATP is formed.• It does not utilize oxygen (anaerobic).

Preparatory (prep) reaction• Both molecules of pyruvate are oxidized and enter

the matrix of the mitochondria.• Electron energy is stored in NADH.• Two carbons are released as CO2 (one from each

pyruvate).

7

Phases of Cellular Respiration Citric acid cycle

• Occurs in the matrix of the mitochondrion and produces NADH and FADH2

• A series of reactions, releases 4 carbons as CO2• Turns twice per glucose molecule (once for each pyruvate)• Produces two immediate ATP molecules per glucose

molecule Electron transport chain (ETC)

• A series of carriers on the cristae of the mitochondria• Extracts energy from NADH and FADH2• Passes electrons from higher to lower energy states • Produces 32 or 34 molecules of ATP by chemiosmosis

8

The Four Phases of Complete Glucose Breakdown

9


Electron transportchain and

chemiosmosis

Mitochondrion

4 ADP

2

4 ATP total

net gain

2 ATP

Glycolysis

NADH

glucose pyruvate

Cytoplasm

ATP

e–

e–e–e–

e–

2 ATP


10

Preparatory reaction

NADH

NADH andFADH2


chemiosmosis

Mitochondrion

2

4 ATP total

net gain

2 ATP

Glycolysis

NADH

glucose pyruvate

Cytoplasm

ATP

e–

e–

e–

2 ATP

e–

e–

e–

4 ADP



11

Mitochondrion

Citric acid cycle


2 ADP 2

NADH

Glycolysis

NADH

glucose pyruvate

Cytoplasm

ATP2

4 ATP total

net gain

2 ATP

ATP

2 ATP

4 ADP


chemiosmosis

NADH andFADH2

e–

e–

e–

e–e–

e–

e–



12

Mitochondrion


2 32 ADPor 34

32or 34

NADH

Glycolysis

NADH

glucose pyruvate

Cytoplasm

ATP

e–

e–

e–

e–

e–

e–

e–

Citric acid cycle


chemiosmosis

NADH andFADH2

2

4 ATP total

net gain

2 ATP

ATP

2 ATP

4 ADP

ATP2 ADP


8.2 Outside the Mitochondria: Glycolysis

• Glycolysis occurs in cytoplasm outside mitochondria.• It consists of a series of 10 reactions, each with its own

enzyme.• Energy Investment Step:

Two ATP are used to activate glucose. Glucose splits into two G3P molecules.

• Energy Harvesting Step: Oxidation of G3P occurs by removal of electrons and hydrogen

ions. Two electrons and one hydrogen ion are accepted by NAD+

resulting in two NADH. Four ATP are produced by substrate-level phosphorylation.

• An enzyme passes a high-energy phosphate to ADP, making ATP. There is a net gain of two ATP (4 ATP produced - 2 ATP

consumed). Both G3Ps convert to pyruvates. 13

Outside the Mitochondria: Glycolysis

14


2

2

4

Glycolysisinputs6C glucose2 NAD+

ATP

4 ADP + 4 P

ATP net gain

totalATP

2 ADP

2 NADHpyruvate

outputs2 (3C)

Glycolysis


G3P glyceraldehyde-3-phosphate

BPG 1,3-bisphosphoglycerate

3PG 3-phosphoglycerate

Energy-investment Step

glucose

Glycolysis

Glycolysis





ADPADP

ATP ATPglucose

Glycolysis


Glycolysis





Two ATP are used to get started.

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

Glycolysis






Splitting produces two3-carbon molecules.

G3PG3P

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

GlycolysisCopyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.






Oxidation of G3P occurs asNAD+ receives high-energyelectrons.

Energy-harvesting Steps

NADH NADH

BPG BPG

G3PG3PNAD+NAD+

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

E1


ADP







Substrate-level ATP synthesis.

+2 ATP ATP

3PG


NADH NADH

ATP

BPG BPGADP

G3PG3P

NAD+NAD+

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

3PG

E2

E1


ADP








Oxidation of 3PG occurs byremoval of water.

+2 ATP ATP

3PG

PEP PEP


NADH NADH

ATP

BPG BPGADP

G3PG3PNAD+NAD+

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

3PG

E3

E2

E1

H2O H2O


ADP

2








Oxidation of 3PG occurs byremoval of water.


Two molecules of pyruvateare the end products of glycolysis.

pyruvatepyruvateATP

+2 ATP

(net gain)

ATP

+2 ATP ATP

3PG

PEP PEPADPADP


NADH NADH

ATP

ATP

BPG BPG

ADP

G3PG3P

NAD+NAD+

ADPADP

–2 ATP


ATP ATP

glucose

Glycolysis

3PG

E4

E3

E2

E1

H2O H2O

Substrate-Level ATP Synthesis

23


enzyme

ADP

ATP

BPG

3PG

8.3 Outside the Mitochondria: Fermentation

• Pyruvate is a pivotal metabolite in cellular respiration.

• If O2 is not available to the cell, fermentation, an anaerobic process, occurs in the cytoplasm. During fermentation, glucose is incompletely

metabolized to lactate, or to CO2 and alcohol (depending on the organism).

• If O2 is available to the cell, pyruvate enters the mitochondria for aerobic respiration. 24

Outside the Mitochondria: Fermentation

• Fermentation is an anaerobic process that reduces pyruvate to either lactate or alcohol and CO2.

• NADH transfers its electrons to pyruvate. • Alcoholic fermentation, carried out by yeasts, produces

carbon dioxide and ethyl alcohol. Used in the production of alcoholic spirits and breads

• Lactic acid fermentation, carried out by certain bacteria and fungi, produces lactic acid (lactate). Used commercially in the production of cheese, yogurt, and

sauerkraut.• Other bacteria produce chemicals anaerobically,

including isopropanol, butyric acid, proprionic acid, and acetic acid.

25

Fermentation

26

2

4

2

glucose

– 2 ATP ATP

E1

G3P

2NAD;

2 NADH

Plants

2CO 2

Animals

2 alcoholor2 lactate

ATP

ATP+ 4

2ADP

E2

BPG4ADP

E3

pyruvate

(net gain)

E4

ATP


or

Fermentation and Food Production

27


© McGraw Hill Education./John Thoeming, photographer


28© Bruce M. Johnson


29© Bruce M. Johnson


• Advantages Provides a quick burst of ATP energy for muscular activity.

• When muscles are working vigorously for a short period of time, lactic acid fermentation provides ATP.

• Disadvantages Lactate and alcohol are toxic to cells. Lactate changes pH and causes muscles to fatigue.

• Oxygen debt Yeast die from the alcohol they produce by fermentation.

• Efficiency of Fermentation Two ATP produced per glucose of molecule during fermentation

is equivalent to 14.6 kcal. Complete oxidation of glucose can yield 686 kcal.

• Efficiency is 2.1% of total possible for glucose breakdown. Only 2 ATP per glucose are produced, compared to 36 or 38

ATP molecules per glucose produced by cellular respiration. 30


31

Fermentationinputs outputs

2 lactate or2 alcohol and 2 CO2

glucose

2 ADP + 2 P net gainATP2


8.4 Inside the Mitochondria• The preparatory (prep) reaction

It connects glycolysis to the citric acid cycle.

End product of glycolysis, pyruvate, enters the mitochondrial matrix.

Pyruvate is converted to a 2-carbon acetyl group.

• Attached to Coenzyme A to form acetyl-CoA

• Electrons picked up (as hydrogen atom) by NAD+, producing NADH

• CO2 released and transported out of mitochondria into the cytoplasm

• Occurs twice per glucose molecule32

Inside the Mitochondria

33

2 NAD+ 2 NADH

2 2+ 2 CoA + 2 CO2

2 pyruvate

O OHC

CH3

+ 2 CoA

CoA

CH3pyruvate

carbondioxideacetyl CoA

C OC O

2 acetyl CoA + 2 carbondioxide


Mitochondrion Structure and Function

34


• Citric Acid Cycle Also called the Krebs cycle Occurs in the matrix of mitochondria Begins with the addition of a 2-carbon acetyl group

(from acetyl-CoA) to a 4-carbon molecule (oxaloacetate), forming a 6-carbon molecule (citric acid)

NADH and FADH2 capture energy rich electrons ATP formed by substrate-level phosphorylation Turns twice for one glucose molecule (once for

each pyruvate) Produces 4 CO2, 2 ATP, 6 NADH, and 2 FADH2 per

glucose molecule

35

Citric Acid CycleCopyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

2

e–

e–

1. The cycle begins when a C2 acetyl group carried byCoA combines with a C4molecule to form citrate.

acetyl CoAC2

oxaloacetateC2

CoA

Glycolysis

glucose pyruvatePreparatory reaction

Electron transportchain andchemiosmosis

Matrix

Citric acidcycle

NADH andFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total

2ADP 2 ATP 32ADPor 34

32or 34

ATP

NADHNADH


2

e–

e–


acetyl CoAC2

oxaloacetateC2

citrateC6

CoA

Glycolysis



Matrix

Citric acidcycle

NADHandFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total


32or 34

ATP

NADHNADH


2

e–

e–

2. Twice over, substratesare oxidized as NAD+ isReduced to NADH,and CO2 is released.


acetyl CoAC2

oxaloacetateC2

citrateC6

NAD

NADH

CoA

ketoglutarateC5

CO2

Glycolysis



Matrix

Citric acidcycle

NADHandFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total


32or 34

ATP

NADHNADH

Citric Acid Cycle

2

e–

e–

3. ATP is produced as anenergized phosphate istransferred from a substrate to ADP.



acetyl CoAC2

oxaloacetateC2

citrateC6

NAD

ATP

succinateC4

NADH

CoA

ketoglutarateC5

NAD+

NADHCO2

CO2

Glycolysis



Matrix

Citric acidcycle

NADHandFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total


32or 34

ATP

NADHNADH


Citric Acid Cycle

2

e–

e–




4. Again a substrate isoxidized, but this timeFAD is reduced to FADH2

acetyl CoAC2

oxaloacetateC2

citrateC6

NAD

fumarateC4

FADH

ATP

succinateC4

NADH

CoA

ketoglutarateC5

NAD+

NADHCO2

FAD

CO2

Glycolysis



Matrix

Citric acidcycle

NADHandFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total


32or 34

ATP

NADHNADH


Citric Acid Cycle

2

e–

e–




5. Once again a substrateis oxidized, and NAD+

is reduced to NADH.

4. Again a substrate isoxidized, but this timeFAD is reduced to FADH2

acetyl CoAC2

NADHoxaloacetate

C2

citrateC6

NAD

fumarateC4

NAD+

FADHATP

succinateC4

Citric acidcycle

NADH

CoA

ketoglutarateC5

NAD+

NADHCO2

FAD

CO2

Glycolysis



Matrix

Citric acidcycle

NADHandFADH2

e–

e–

e–

e–

e–

2 ATP

2 ADP

netATP

4 ADP 4 ATP total


32or 34

ATP

NADHNADH



42

inputs outputs4 CO2

NADH

FADH22

2 (2c) acetyl groups

6 NAD+

2 FAD

2

Citric acid cycle

P2 ADP +2

6

ATP


Inside the Mitochondria• Electron Transport Chain (ETC)• Location:

Eukaryotes – cristae of the mitochondria Aerobic prokaryotes – plasma membrane

• Series of carrier molecules: Pass energy-rich electrons successively from one to another Complex arrays of protein and cytochrome

• Proteins with heme groups with central iron atoms

• The electron transport chain: Receives electrons from NADH & FADH2 Produces ATP by oxidative phosphorylation

• Oxygen final electron acceptor: Combines with hydrogen ions to form water

43

Inside the Mitochondria• The fate of the hydrogens Hydrogens from NADH deliver enough energy to

make 3 ATPs. Those from FADH2 have only enough for 2 ATPs. “Spent” hydrogens combine with oxygen.

• Recycling of coenzymes increases efficiency. Once NADH delivers hydrogens, it returns (as NAD+)

to pick up more hydrogens. However, hydrogens must be combined with oxygen

to make water. If O2 is not present, NADH cannot release H+. It is no longer recycled back to NAD+.

44

45

Inside the Mitochondria• The electron transport chain complexes pump H+ from the

matrix into the intermembrane space of the mitochondrion.

• H+ therefore becomes more concentrated in the intermembrane space, creating an electrochemical gradient.

• ATP synthase allows H+ to flow down its gradient.• The flow of H+ drives the synthesis of ATP from ADP and

inorganic phosphate by ATP synthase.• This process is called chemiosmosis.

ATP production is linked to the establishment of the H+ gradient.• ATP moves out of mitochondria and is used for cellular

work. It can be broken down to ADP and inorganic phosphate. These molecules are returned to the mitochondria for more ATP

production.

Organization and Function of the Electron Transport Chain


2

O212+

2

P

H+

e:

e :

Electron transport chain

high energyelectron

ATPchannelprotein

Intermembranespace

ATPsynthasecomplex

low energyelectron

cytochromeoxidase

cytochrome c

NADH-Qreductase

cytochromereductase

coenzyme Q

NADH

Matrix

FADH2 FAD+

NAD+

2

e-

2 ADP ATP32 ADPor 34

4 ADP

netATP

4 ATP total

2ADP

2ATP

Glycolysis

glucose pyruvatePreparatory reaction Citric acid

cycle


chemiosmosis

NADH andFADH2

NADH

NADHe-

e-

e-e-

e-

e-

32 or34

H+

H+

H+

H+

H+

H+

H+

H+

H+

H+

H+

H+

H+H+H+

H+

H+

H+

ATP ADPH2O

H+

H+

H+

H+

H+

H+

H+

H+

H+H+H+

H+

H+

H+H+

H+

H+ATP

Chemiosmosis


• Energy yield from glucose metabolism Net yield per glucose

• From glycolysis – 2 ATP• From citric acid cycle – 2 ATP• From electron transport chain – 32 or 34 ATP

Energy content• Reactant (glucose) 686 kcal• Energy yield (36 ATP) 263 kcal• Efficiency is 39%• Rest of energy from glucose is lost as heat

47

Energy Yield from Glucose Metabolism

48


glucose


49

Cyt

opla

sm

Elec

tron

tran

spor

t cha

in

2net

glucose

2 pyruvate

glycolysis

NADH2 4 or 6


ATP

ATP


50


Cyt

opla

sm

Elec

tron

tran

spor

t cha

in

2net

glucose

2 pyruvate

2 acetyl CoA

glycolysis

NADH

NADH

2

2

4 or 6

6

ATP

ATP

ATP


51

Cyt

opla

sm

Elec

tron

tran

spor

t cha

in

2net

2

glucose

2 pyruvate

2 acetyl CoA

Citric acidcycle

glycolysis

2 CO2

NADH

NADH

NADH

FADH2

2

2

6

2

4 or 6

6

18

4


ATP

ATP

ATP

ATP

ATP

4CO2

6O2

ATP

6HO2


52

Cyt

opla

sm

Elec

tron

tran

spor

t cha

in

2net

2

glucose

2 pyruvate

2 acetyl CoA

subtotal subtotal

glycolysis

NADH

NADH

NADH

2

2

6

2

4 or 6

6

18

4

324or 34


ATP

ATP

ATP

4CO2

2CO2

FADH2

6O2 6HO2

ATP

ATP

ATP

ATP

ATP

Citric acidcycle


53

Cyt

opla

smM

itoch

ondr

ion

Elec

tron

tran

spor

t cha

in

2net

2

glucose

2 pyruvate

2 acetyl CoA

Citric acidcycle

subtotal subtotal

glycolysis

2 CO2

4 CO2

NADH

NADH

NADH

FADH2

2

2

6

2

4 or 6

6

18

4

324

36 or 38total

6 O2 6 H2O

ATP

ATP

ATP

ATP

ATP

ATPATP

ATP

ATPor 34


8.5 Metabolism• Foods Sources of energy rich molecules

Carbohydrates, fats, and proteins

• Degradative reactions (catabolism) break down molecules. Tend to be exergonic (release energy)

• Synthetic reactions (anabolism) build molecules. Tend to be endergonic (consume energy)

54

The Metabolic Pool Concept

55

ATP

ATPElectrontransportchain

proteins carbohydrates fats

aminoacids

glucose fattyacids

glycerol

acetyl CoA

Glycolysis

pyruvate

ATPCitricacidcycle


© C Squared Studios/Getty RF

Metabolic Pool • Glucose is broken down in cellular respiration.• Fat breaks down into glycerol and three fatty

acids.• Amino acids break down into carbon chains and

amino groups. Deamination (NH2 removed) occurs in the liver.

• Results in poisonous ammonia (NH3)• Quickly converted to urea

Different R-groups from amino acids are processed differently.

Fragments enter respiratory pathways at many different points.

56

Metabolic Pool• All metabolic compounds are part of the

metabolic pool.• Intermediates from respiratory pathways can be

used for anabolism.• Anabolism (synthetic reactions of metabolism): Carbohydrates

• Start with acetyl-CoA• Basically reverses glycolysis (but different pathway)

Fats• G3P converted to glycerol• Acetyl groups are connected in pairs to form fatty acids.

57

Metabolic Pool• Anabolism (cont’d): Proteins

• They are made up of combinations of 20 different amino acids.

• Some amino acids (11) can be synthesized by adult humans.

• However, other amino acids (9) cannot be synthesized by humans.

– Essential amino acids– Must be present in the diet

58

The Energy Organelles Revisited• Similarities between photosynthesis and cellular

respiration: Use of membrane

• Chloroplasts’ inner membrane forms thylakoids.• Mitochondria’s inner membranes form cristae.

Electron transport chain ETC is located on thylakoid membranes and cristae.

In photosynthesis, electrons passed to ETC were energized by the sun.

In mitochondria electrons, energized electrons were removed from glucose.

In both, ETC establishes an electrochemical gradient of H+ with ATP production by chemiosmosis.

Enzymes• In chloroplast, stroma has Calvin cycle enzymes.• In mitochondria, matrix contains enzymes of citric acid cycle. 59

Flow of Energy• Energy flows from the sun through chloroplasts to

carbohydrates and then through mitochondria to ATP molecules. This flow of energy maintains biological organization at all

levels from molecules to organisms to the biosphere. Some energy is lost with each chemical transformation.

• Eventually all solar energy captured is lost.• All life depends on solar energy input.

• Chemicals cycle within natural systems. Chloroplasts produce oxygen and carbohydrates, which are

used by mitochondria to generate energy for life.• Chloroplasts and mitochondria allow energy flow

through organisms and permit chemical cycling.60

The Energy Organelles Revisited

61


O2

O2

enzymesgranaPhotosynthesis

NADP+

CH2O

NADPH

CO2

enzyme-catalyzedreactions

NADH

enzymes

H2O

membrane

ATP productionvia chemiosmosis

ADP ATP

H2O

membrane

Cellular RespirationNAD+

cristaeCH2O CO2

biology - pccua...respiration. • if o 2 is not available to the cell, fermentation, an anaerobic...

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